Iowa State University

From the SelectedWorks of Kevin J. Roe

2005

Hamiota, a new genus of freshwater mussel (: ) from the Gulf of Mexico drainages of the southeastern United States Kevin J. Roe Paul D. Hartfield

Available at: https://works.bepress.com/kevin_roe/9/ THE NAUTILUS 119(1):1–10, 2005 Page 1

Hamiota, a new genus of freshwater mussel (Bivalvia: Unionidae) from the Gulf of Mexico drainages of the southeastern United States

Kevin J. Roe Paul D. Hartfield Delaware Museum of Natural History U.S. Fish and Wildlife Service 4840 Kennett Pike 6578 Dogwood View Parkway, Suite A Wilmington, DE 19807 USA Jackson, MS 39213 USA [email protected] Paul࿞Hartfi[email protected]

ABSTRACT MATERIALS AND METHODS

Hamiota, a new genus of freshwater mussel containing four A list of specimens examined is included in Appendix 1. formerly assigned to the genus Lampsilis Rafinesque, Acronyms used in the text are: Academy of Natural Sci- 1820, is described. In addition to the genus Lampsilis, mem- ences of Philadelphia (ANSP), Delaware Museum of bers of Hamiota had previously been placed in the genera Vil- Natural History (DMNH), Florida Museum of Natural losa Frierson, 1927, and Ligumia Swainson, 1840. Several char- History (UF), United States National Museum (USNM), acters including the packaging of their larvae in a superconglu- University of Alabama Unionid Collection (UAUC), and tinate lure to attract host fishes, placement and shape of the Museum of Natural Science (MMNS). Ab- marsupia, and release of glochidia through the excurrent si- breviated synonymies are presented for each taxon and phon, support the recognition of these species as a distinct include novel combinations and publications with illus- genus. tration. Measurements were taken to the nearest 0.05 mm using dial calipers.

SYSTEMATICS INTRODUCTION Family Unionidae Rafinesque, 1820 Following the discovery in 1988 by Robert Butler of the Tribe Lampsilini von Ihering, 1901 first superconglutinate lure ensnared on a snag in a trib- Hamiota new genus. Type species: Hamiota subangulata (Lea, utary of the Choctawhatchee River, a number of publi- 1840) by original designation cations (e.g., Haag et al., 1995; Hartfield and Butler, 1997; O’Brien and Brim Box, 1999; Blalock-Herod et al., Diagnosis: A monophyletic group of freshwater bi- 2002) have confirmed through direct observation the valves (Roe et al., 2001) in which all of the glochidia are supposition that these unique lures are produced by four released simultaneously encased in mucous packages species of freshwater mussels endemic to the Gulf of that are referred to as superconglutinates (Haag et al., Mexico drainages of the southeastern United States. 1995, fig. 1). The superconglutinate lure exits the mantle Herein, we confirm earlier published suggestions that cavity via the excurrent opening and is encased within a these four species represent a distinct genus of fresh- transparent mucous tube (Hartfield and Butler, 1997; water mussels (Fuller and Bereza, 1973; O’Brien and O’Brien and Brim Box, 1999). When acted upon by wa- Brim Box, 1999). The recognition of this genus is based ter currents the superconglutinate mimics the move- on several characters including the production of the su- ments of a swimming fish, and has been shown to elicit perconglutinate lure, and the unique shape and place- attacks from fishes (Haag and Warren, 1999). The mar- ment of the marsupia (the region of the demibranchs supium is restricted to the ventral portion of the outer where female unionoid mussels brood developing lar- demibranchs of female mussels (Figure 1). The precise vae), and is supported by molecular evidence (Roe et al., shape and pigmentation of the marsupia, as well as the 2001). Use of marsupial features is consistent with pre- degree of posterior mantle margin development, varies vious designations of unionid ‘‘divisions’’ initiated by across species. Simpson (1900a) and continued by Heard and Guckert Description: Members of this genus are small- to me- (1970). The designation of Hamiota increases the num- dium-sized freshwater bivalves, and adult valves gener- ber of North American unionids genera to 50 (Turgeon ally are between 45–100 mm in length. Shells range et al., 1998). from ovate to elliptical in outline, and are somewhat Page 2 THE NAUTILUS, Vol. 119, No. 1

Figures 1, 2. Inner mantle and outer demibranchs of gravid superconglutinate-producing mussels in the genus Hamiota and inner mantle and outer demibranchs typical of non-superconglutinate-producing mussels of the genera Lampsilis, Ligumia, and Villosa. 1. Female Hamiota australis. 2. Female Villosa vibex. compressed to moderately inflated. Shell thickness rang- and colors may include purple, red, black, or white. In es from heavy to thin. Sexes display some degree of di- females, the mantle margins anterior to the branchial morphism in shell shape. Shells of male mussels are typ- opening are elaborated to varying degrees. The mor- ically more acutely pointed posteriorly, whereas shells of phology of the glochidial valves of members of Hamiota female mussels display an expanded posterior margin. is similar to that of members of Lampsilis or Villosa Periostracum is typically smooth, but can be very glossy (O’Brien and Brim Box, 1999). in some species. Background color ranges from dark- Etymology: Hamiota ϭ angler. Derived from the brown and black through chestnut-brown to straw-yel- Greek word hamus, meaning hook. This name refers to low. Black to bright green rays of variable width are of- the means by which members of this genus attract host ten present and may be limited to the posterior slope, fishes by packaging their parasitic larvae in a lure that or cover the entire disk. Nacre color is typically white mimics a small fish. although other colors such as salmon or blue may be seen as well, particularly in the beak cavities or the pos- Remarks: Species of Hamiota generally have been terior margins. The marsupia are often asymmetrical in treated as Lampsilis due to the similarity in shape and shape; the anterior portion is typically broadest, tapering coloration of their shells. Some authors also have in- toward the posterior end. The ventral margin of the mar- cluded these species in the genus Villosa or Ligumia, supium is darkly pigmented in gravid females. Pigmen- due to shell shape, thickness, and/or ornamentation of tation of marsupia varies across species and populations the mantle flap. The shells of Hamiota species are in- K. J. Roe and P. D. Hartfield, 2005 Page 3 deed similar to the shells of these genera, and the man- mantle flap is typically not expanded and is reddish in tle flap may exhibit characters of both Lampsilis and color. Marsupia of H. altilis are finely tapered at each Villosa. However, in Hamiota, the marsupia is restricted end when immature, becoming broadly rounded on the to the ventral half of the posterior portion of the outer ends in most populations, tapering anteriorly in others. demibranchs, while in Lampsilis, Ligumia, and Villosa Marsupia color is a dark reddish-brown or black along the marsupium fully occupies the water tubes of the pos- the margin and white above. The anus is usually pig- terior portion of the outer demibranchs (Figure 1). Un- mented red and the incurrent and excurrent siphons are like members of Lampsilis, which release larvae through reddish-brown to black. The glochidia of H. altilis are pores in the ventral edge of the demibranchs, members described by Haag et al. (1999). of Hamiota release the superconglutinate lure through Type Material: Unio altilis Conrad, 1834, Lectotype the excurrent opening. The placement and shape of the ANSP 56419 (Figures 3, 4) here designated. Type lo- marsupium and the extraordinary method of glochidial cality: Alabama River, near Claiborne [Monroe Co., Al- release and host fish attraction, the superconglutinate, abama]. are uniquely derived characters in the Lampsilini that Unio clarkianus Lea, 1852, Type not found. Type local- warrant genus-level recognition. ity: Williamsport, [Maury Co.], Tennessee; Georgia or Alabama. (Conrad, 1834) new combination Unio doliaris Lea, 1865, Lectotype USNM 84936, here designated. Type locality: Etowah River, Georgia. Unio altilis Conrad, 1834; Conrad, 1834: 43, pl. 2, fig. 1; Unio gerhardtii Lea, 1862, Holotype USNM 25711 by Chenu, 1845: 21, pl. 1, fig. 1; Reeve, 1865: pl. 23, fig. 109. monotypy. Type locality: Chattanooga, Georgia. Margarita (Unio) altilis (Conrad, 1834); Lea, 1836: 24. Margaron (Unio) altilis (Conrad, 1834); Lea, 1852a: 27. Remarks: The most variable species included in Ham- Lampsilis altilis (Conrad, 1834); Simpson, 1900a: 529; Par- iota, H. altilis, is endemic to the Mobile River Basin. malee and Bogan, 1998: 125, pl. 47. Some of the conchological variation is undoubtedly eco- Unio clarkianus Lea, 1852; Lea, 1852b: 251; Lea, 1852c: 273, phenotypic in nature, although the extent and nature of pl. 21, fig. 30; Lea, 1852d: 29, pl. 21, fig. 30. Margaron (Unio) clarkianus (Lea, 1852); Lea, 1852a: 27. the variation in shell shape and pigmentation has not Lampsilis clarkianus (Lea, 1852); Simpson, 1900a: 532. been adequately explored. Unio gerhardtii Lea, 1862; Lea, 1862a: 168; Lea, 1862b: 208, Conrad (1834) in his original description did not iden- pl. 31, fig. 277; Lea, 1862c: 30, pl. 31, fig. 277. tify a primary type. Johnson and Baker (1973) identified Margaron (Unio) gerhardtii (Lea, 1862); Lea, 1870: 35. ANSP 56419 as the figured holotype, although the spec- Lampsilis (Lampsilis) gerhardtii (Lea, 1862); Simpson, 1900a: imen label indicates the locality as ‘‘Ogeechee R., Ga.’’ 532. Johnson and Baker (1973) state that the label is in error, Unio doliaris Lea, 1865; Lea, 1865: 88; Lea, 1868: 260, pl. 32, and ‘‘probably was mixed with ANSP 46418, which is fig. 75; Lea, 1869: 20, pl. 32, fig. 75. labeled ‘Claiborne, Alabama’, by error.’’ Conrad (1834) Margaron (Unio) doliaris (Lea, 1865); Lea, 1870: 42. clearly indicated that the specimen(s) of U. altilis de- Lampsilis (Lampsilis) doliaris (Lea, 1865); Simpson, 1900a: 533. scribed were collected from the ‘‘Alabama River, near Claiborne.’’ Conrad (1834) did not specifically designate Description: Described by Conrad (1834) as sub-oval, a holotype and according to the ICZN recommendation thin, and inflated. The periostracum was described as 73F and Article 74.5 the holotype designation of John- ‘‘rugose’’ and ‘‘blackish’’ with rays and ‘‘numerous short son and Baker (1973) is deemed invalid. In order to vermicular lines on the posterior slope,’’ and the nacre preserve stability of nomenclature, we herein designate as whitish and iridescent. The periostracum of speci- specimen ANSP 56419 as the lectotype of U. altilis. In mens of H. altilis is typically brown to chestnut-brown accordance with ICZN Article 74.7, we herein designate in color with a variable number of dark green rays. The the specimen USNM 84936 as the lectotype of U. do- left valve has two heavy, spatulate pseudocardinal teeth, liaris Lea in order to maintain taxonomic stability and the smaller above the larger. The right valve has two because this specimen appears to be that figured by Lea nearly equally sized triangular teeth, the larger anterior (1868). Hamiota altilis is considered threatened by the to the smaller. The lateral teeth are short but blade-like, United States Fish and Wildlife Service (USFWS, 1994). two in the right valve, and one in the left. Life History: Mature gravid females have been re- The posterior mantle margin of the female is expand- ported from March through June. Hamiota altilis have ed into a well-developed flap with papillae along the bor- also been observed releasing glochidia in a superconglu- der. Coloration, number, and size of papillae vary some- tinate (Haag et al., 1999). Large centrarchid fishes, in- what between populations. In general, however, the in- cluding Micropterus coosae Hubbs and Bailey, 1940, M. terior mantle flap is colored red to dark red or brown punctulatus (Rafinesque, 1819), M. salmoides (Lace- with darker spots, while the exterior of the flap is brown pe´de, 1802), and Lepomis cyanellus Rafinesque, 1819, to black, often with vertical lighter bars, and with a have been confirmed as suitable hosts (Haag et al., small, but prominent dark ‘‘eye spot’’ on the posterior 1999). end. Small papillae are present along the mantle flap, usually becoming more robust anteriorly. In males, the Range: Hamiota altilis was historically reported Page 4 THE NAUTILUS, Vol. 119, No. 1

Figures 3–10. Type material of species of Hamiota. Photographs are of the interior of left valve and exterior of right valve. 3, 4. Lectotype of H. altilis ANSP 56419. 5, 6. Holotype of H. australis USNM 150473. 7, 8. Lectotype of H. perovalis ANSP 56416. 9, 10. Lectotype of H. subangulata USNM 85801. K. J. Roe and P. D. Hartfield, 2005 Page 5 throughout the Mobile River Basin, including the Tom- cause both were described from practically the same lo- bigbee, Black Warrior, Cahaba, Alabama, Tallapoosa, and cality. An examination of mitochondrial DNA sequences Coosa River drainages in Alabama, Georgia, Mississippi, by Roe et al. (2001) recovered these two taxa as a clade, and Tennessee. The species is currently restricted to lo- but failed to resolve them into reciprocally monophyletic calized portions of the Cahaba, Coosa, and Tallapoosa groups. Hurd (1974) considered perovalis a junior syn- rivers and some of their tributaries (USFWS, 2003). onym of altilis, as did Burch (1975). Frierson (1927) considered U. doliaris (Lea, 1865) a synonym of perov- alis, although Parmalee and Bogan (1998) include U. do- (Conrad, 1834) new combination liaris as a synonym of altilis. Based on the collection Unio perovalis Conrad, 1834; Conrad, 1834: 43, pl. 2, fig. 2; locality and the appearance of the type specimen, we Chenu, 1845: 21, pl. 1, fig. 2; Ku¨ ster, 1861: 257, pl. 87, place U. spillmanii Lea, 1861 as a synonym of H. per- fig. 2; Reeve, 1866: pl. 38, fig. 209. ovalis. As with U. altilis, Conrad (1834) did not specifi- Margarita (Unio) perovalis (Conrad, 1834); Lea, 1836: 24. cally designate a holotype for U. perovalis, therefore, Margaron (Unio) perovalis (Conrad, 1834); Lea, 1852a: 27. according to the ICZN Article 74.5 and recommendation Lampsilis perovalis (Conrad, 1834); Simpson, 1900a: 531. 73F the holotype designation of Johnson and Baker Unio spillmanii Lea, 1861; Lea, 1861: 39, Lea, 1862d: 98, pl. (1973) is deemed to be in error. In an effort to maintain 15, fig. 246; Lea, 1862e: 102, pl. 15, fig. 246, Reeve, 1868: nomenclatural stability we herein designate ANSP 56419 pl. 82, fig. 435. Margaron (Unio) spillmanii (Lea, 1861); Lea, 1870: 42. the lectotype for U. perovalis. According to ICZN Arti- Lampsilis (Lampsilis) spillmani (Lea, 1861); Frierson, 1927: 69 cle 74.7, the specimen USNM 84925 is here designated [misspelling]. as the lectotype of U. spilmanii in order to fix the name and maintain nomenclatural stability. This species is list- Description: Conrad (1834) described this species as ed as threatened by the United States Fish and Wildlife oval and inflated with a moderately thick shell. He noted Service (USFWS, 1994). two color varieties, one in which the periostracum was olivaceous and obscurely rayed with white nacre and an- Life History: Discharge of superconglutinates was other in which the periostracum was reddish-brown with first observed in H. perovalis (Haag et al., 1995). Glo- ‘‘rose colored’’ nacre. The periostracum of specimens of chidia mature and are discharged between March and H. perovalis is generally lighter in color than H. altilis June, with releases concentrated in early April (Hartfield and range from straw-yellow to light brown. The number and Butler, 1997). Micropterus coosae, M. punctulatus, of rays is variable and can cover the entire disk. The left and M. salmoides have been identified as suitable host valve contains two robust equal sized pseudocardinal fishes for the orange-nacre mucket (Haag and Warren, teeth. The right valve has two pseudocardinals and the 1999). anterior tooth is smaller than the posterior tooth. The Range: Hamiota perovalis was historically known from lateral teeth are elongate, two in the left, one in the the Mobile Basin’s Alabama, Tombigbee, Black Warrior, right. and Cahaba rivers and their tributaries in Alabama and The mantle margins of female H. perovalis are ex- Mississippi. The species has apparently become extir- panded into well-developed flaps, pigmented red on the pated from the main channels of the larger rivers, but interior and darker red to brown or black on the exterior. continues to survive in some tributaries of all four drain- No eyespot is present and short papillae are present ages (USFWS, 2003). along the mantle edge. Males possess a rudimentary mantle margin with weak pigmentation and few papillae. The marsupia of H. perovalis are pisciform in shape, Hamiota subangulata (Lea, 1840) new combination broader anteriorly and narrowly tapering posteriorly. The marsupium is reddish or darker along the margin, Unio subangulatus Lea, 1840; Lea, 1840: 287; Lea, 1842a: 209, often with a darker spot of pigment on the broader an- pl. 13, fig. 23; Lea, 1842b: 47, pl. 8, fig. 23; Ku¨ ster, 1861: 278, pl. 94, fig. 2; Simpson, 1892: 415, pl. 58, fig. 1; Reeve, terior end that resembles an eyespot in the supercon- 1868: pl. 65, fig. 327. glutinate lure. The anus can be pigmented red and Margaron (Unio) subangulatus (Lea, 1840); Lea, 1852a: 29. black, and the incurrent and excurrent siphons are usu- Lampsilis subangulatus (Lea, 1840); Simpson, 1900a: 556; ally reddish or brown in color. Clench and Turner, 1956: 196, pl. 2, fig. 2. Ligumia subangulata (Lea, 1840); Haas, 1969: 443. Type Material: Unio perovalis Conrad, 1834, Lecto- Villosa subangulata (Lea, 1840); Heard, 1979: 44. type ANSP 56416 (Figures 7, 8), here designated. Type Unio kirklandianus S. H. Wright, 1897; S. H. Wright, 1897: locality: Alabama River, at Claiborne [Monroe Co., Al- 136. abama]. Lampsilis kirklandianus (S. H. Wright, 1897); Simpson, 1900a: Unio spillmanii Lea, 1861, Lectotype USNM 84925, 557; Simpson, 1900b: 76, pl. 1, fig. 7. here designated. Type locality: Luxpalila Creek, near Co- lumbus, Mississippi. Description: A medium-sized mussel that reaches ap- proximately 85 mm in length (Brim Box and Williams, Remarks: There has been some question as to the dis- 2000). Specimens are generally elongate; the posterior tinctiveness of H. perovalis from H. altilis, perhaps be- ridge is rounded and the posterior slope is usually con- Page 6 THE NAUTILUS, Vol. 119, No. 1 cave. Periostracum color is variable in this species. In- Hamiota australis (Simpson, 1900) new combination dividuals range from straw-yellow to chestnut-brown in color with a variable number of black to bright emerald Lampsilis australis Simpson, 1900; Simpson, 1900a: 544; Simp- son, 1900b: 75, pl. 2, fig. 2; Clench and Turner, 1956: 199, green rays of variable width. These rays emanate from pl. 2, fig. 3. the umbo and radiate across the disk. Most shells are Ligumia australis (Simpson, 1900); Haas, 1969: 432. shiny, although some populations exhibit an extremely Villosa australis (Simpson, 1900); Heard, 1979: 44. glossy periostracum. This species has been described as one of the most beautiful of all North American fresh- Description: Simpson (1900b) described the shells of water mussels (S. H. Wright, 1897; Clench and Turner, this species as ‘‘long and elliptical . . . moderately inflat- 1956). ed.’’ The periostracum was described as ‘‘smooth, shin- Wright (1897) described U. kirklandianus from the ning, greenish yellow, rayed with green’’ and the nacre Ochlockonee River in Leon County, Florida and re- as ‘‘bright bluish white.’’ Specimens examined were el- marked that his specimens were more polished and had liptical to elongate oval and often terminate in a blunt broader rays than typical of U. subangulatus, and that point. Specimens are almost always stained black, with the shells were ‘‘deeper and broader.’’ The right valve some green rays visible on the posterior slope. The per- has two somewhat spatulate pseudocardinal teeth, the iostracum is often glossy. The right valve contains two smaller nearly directly above the other. The left valve pseudocardinal teeth, the smaller above the larger. The has two pseudocardinal teeth, the anterior one much left valve has two compressed pseudocardinal teeth. Lat- larger than the other. Lateral teeth are thin, but not eral teeth are elongate and slightly curved, two in the delicate, two in the left and one in the right valve. Ex- left valve and one in the right. Posterior mantle flaps are amination of specimens from across the range of this poorly developed with a streak of red stain along the species reveals substantial variation in shell color and margins. There is no eyespot on the mantle flap, and size. The mantle margins in females are only slightly ex- only a few very small papillae. The marsupia are pisci- panded into a modest flap. The flaps are light brown in form, broadly rounded anteriorly and tapering behind coloration on the interior and freckled-brown on the ex- (Figure 1). Color of the marsupium is white along the terior, and no eyespot is present. Short papillae are pres- margin and black above. The anus is unpigmented and ent along the margin of the flap, becoming larger ante- the incurrent and excurrent siphons are reddish in color. riorly. In males, the mantle is only slightly expanded, Glochidia were described and figured by Blalock-Herod without pigment, and with very short papillae. The mar- et al. (2002). supia are pisciform in shape, broader anteriorly and ta- Type Material: Lampsilis australis Simpson, 1900, pered behind, and darkly pigmented along the margin Holotype USNM 150473 by original designation (Fig- often with a darker spot of pigment anteriorly. The anus ures 5, 6). Type locality: Little Patsiliga Creek, south- is unpigmented and the siphons are brownish. Glochidia eastern Alabama. were figured and described by O’Brien and Brim Box (1999). Remarks: Fuller and Bereza (1973) stated that this species represented an ‘‘undescribed lampsiline genus’’ Type Material: Unio subangulatus Lea, 1840, Lecto- and that its marsupium allied it closely with Ptychobran- type USNM 85801 (Figures 9, 10) designated by Clench chus. This species was incorrectly synonymized with Pty- and Turner (1956). Type locality: Chattahoochee River, chobranchus jonesi (van der Schalie, 1934) by Clench Columbus, [Muscogee Co.,] Georgia. and Turner (1956). The phylogenetic analysis of Roe et Unio kirklandianus S. H. Wright, 1897, Paratype USNM al. (2001) clearly placed this species with the other su- 149648. Type locality: Oclocknee River, Leon Co., Flor- perconglutinate producers. ida. Life History: Superconglutinate releases have been Remarks: Hamiota subangulata is listed as an endan- documented in this species by Blalock-Herod et al. gered species (USFWS, 1998). The analysis of Roe et al. (2002). Micropterus spp. are likely hosts (Blalock-Herod (2001) resolved this taxon as monophyletic and in a clade et al., 2002). with H. australis. Range: Hamiota australis was known historically from Life History: Discharge of superconglutinate lures the Escambia, Yellow, and Choctawhatchee River sys- has been documented by O’Brien and Brim Box (1999) tems. It continues to survive in some river and stream from late May through early June. Micropterus punc- segments within these systems. This species is not cur- tulatus and M. salmoides appear to be primary hosts for rently protected under the Endangered Species Act. the species (O’Brien and Brim Box, 1999). ACKNOWLEDGMENTS Range: Hamiota subangulata was found throughout the Apalachicola River Basin and the Ochlockonee River We express thanks to the many individuals who have drainage (Brim Box and Williams, 2000). Currently the worked with this unique group of mussels and directly species continues to survive in some small streams and or indirectly assisted in the production of this work. Sev- headwater rivers (USFWS, 1998). eral museum curators provided access to the specimens K. J. Roe and P. D. Hartfield, 2005 Page 7 in their care including the National Museum of Natural Louis, Missouri. Upper Mississippi Conservation Com- History (Dr. Robert Hershler), The Florida Museum of mittee, Rock Island, Illinois, pp. 11–14. Natural History (Dr. Fred Thompson), The Academy of Heard, W. H. 1979. Identification manual of the freshwater Natural Sciences of Philadelphia (Dr. Dan Graf), The clams of Florida. State of Florida Department of Envi- University of Alabama Malacology Collection (Dr. ronmental Regulation, Technical Series, vol.4, no. 2, 83 pp. Charles Lydeard), and the Mississippi Museum of Nat- Heard, W. H. and R. H. Guckert. 1970. A re-evaluation of the ural Science (Dr. Robert Jones and Leann Staton). The recent Unionacea (Pelycepoda) of North America. Mala- comments of several individuals greatly improved this cologia 10: 333–355. manuscript: We are grateful to Art Bogan, Jeffrey Gar- Hubbs, C. L. and R. M. Bailey. 1940. A revision of the black ner, Dan Graf, Jeanne Serb, and Jim Williams. Gary basses (Micropterus and Huro) with descriptions of four Bloomer (DMNH) assisted with figures. new forms. Miscellaneous Publications Museum of Zool- ogy, University of Michigan 48: 1–51. Hurd, J. C. 1974. Systematics and zoogeography of the Union- LITERATURE CITED acean mollusks of the Coosa River Drainage of Alabama, Georgia and Tennessee. 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New fresh water shells of the United descriptions of new species in the families Naı¨ades, Col- States, with coloured illustrations; and a monograph of the imacea, Lymnaeana, Melaniana and Peristomiana. 3: 1– genus Anculotus of Say; also a synopsis of the American 88, pls. 5–27. naiads. Judah Dobson, Philadelphia, 76 pp. ϩ 8 plates. Lea, I. 1852a. A synopsis of the family of Naı¨ades. Philadelphia. Frierson, L. S. 1927. A classified and annotated check list of 3rd edition, Blanchard and Lea, Philadelphia, 88 pp. the North American naiads. Baylor University Press, Lea, I. 1852b. Descriptions of new species of the family Union- Waco, 111 pp. idae. Proceedings of the American Philosophical Society Fuller, S. L. H. and D. J. Bereza. 1973. Recent additions to of Philadelphia 5: 251–252. the naiad fauna of the eastern Gulf Drainage (Bivalvia: Lea, I. 1852c. Descriptions of new species of the family Union- Unionoida: Unionidae). Association of Southeastern Biol- idae. Transactions of the American Philosophical Society ogists Bulletin 20: 53. 10[New Series]: 253–294, pls. 12–29. Haag, W. R., R. S. Butler and P. D. Hartfield. 1995. An ex- Lea, I. 1852d. Observations on the genus Unio, together with traordinary reproductive strategy in freshwater bivalves: descriptions of new species in the families Unionidae, Col- prey mimicry to facilitate larval dispersal. Freshwater Bi- imacea and Melaniana 5: 9–61, pls. 30. ology 34: 471–476. Lea, I. 1861. Descriptions of twenty-five new species of Union- Haag, W. R. and M. L. Warren. 1999. Mantle displays of fresh- idae from Georgia, Alabama, Mississippi, Tennessee and water mussels elicit attacks from fish. Freshwater Biology Florida. Proceedings of the Academy of Natural Sciences 42: 35–40. of Philadelphia 13: 38–41. Haag, W. R., M. L. Warren and M. Shillingsford. 1999. Host Lea, I. 1862a. Descriptions of ten new species of Unionidae fishes and host attracting behavior of Lampsilis altilis and from the United States. Proceedings of the Academy of Villosa vibex (Bivalvia: Unionidae). American Midland Natural Sciences of Philadelphia 14: 168–169. Naturalist 141: 149–157. Lea, I. 1862b. New Unionidae of the United States and arctic Haas, F. 1969. Superfamilia Unionacea. Das Tierreich (Berlin) America. Journal of the Academy of Natural Sciences of 88: x ϩ 663 pp. Philadelphia 5[New Series]: 187–216, pls. 24–33. Hartfield, P. and R. Butler. 1997. Observations on the release Lea, I. 1862c. Observations on the genus Unio, together with of superconglutinates by Lampsilis perovalis (Conrad descriptions of new species, their soft parts, and embry- 1834). In: Cummings, K. S., A. C. Buchanan, C. A. Mayer, onic forms, in the family Unionidae, and descriptions of and T. J. Naimo (eds.) Conservation and management of new genera and species of the Melanidae. 9: 9–178, pls. freshwater mussels II: Initiatives for the future. Proceed- 24–38. ings of a UMRCC symposium, 16–18 October 1995, St. Lea, I. 1862d. New Unionidae of the United States. Journal of Page 8 THE NAUTILUS, Vol. 119, No. 1

the Academy of Natural Sciences of Philadelphia 5 [New status for five freshwater mussels and threatened status Series]: 14–109, pls. 1–18. for two freshwater mussels from eastern Gulf slope drain- Lea, I. 1862e. Observations on the genus Unio, together with ages of Alabama, Florida, and Georgia. Federal Register descriptions of new species, their soft parts, and embry- 63: 12664–12687. onic forms, in the family Unionidae. 8: 9–114, 34 pls. U.S. Fish and Wildlife Service. 2003. Endangered and threat- Lea, I. 1865. Descriptions of eight new species of Unio from ened wildlife and plants; proposed designation of critical the United States. Proceedings of the Academy of Natural habitat for three threatened mussels and eight endangered Sciences of Philadelphia 17: 88–89. mussels in the Mobile River Basin; proposed rule. Federal Lea, I. 1868. New Unionidae, Melanidae, etc., chiefly of the Register 68: 14752–14832. United States. Journal of the Academy of Natural Sciences van der Schalie, H. 1934. Lampsilis jonesi, a new naiad from of Philadelphia 6 [New Series]: 249–302, pls. 29–45. southeastern Alabama. The Nautilus 47: 125–127. Lea, I. 1869. Observations on the genus Unio, together with von Ihering, H. 1901. The Unionidae of North America. The descriptions of new species in the family Unionidae, and Nautilus 15: 37–39, 50–53. descriptions of new species of the Melanidae and Palu- Wright, S. H. 1897. Contribution to the knowledge of the Unit- dinae. 12: 9–105, 26 pls. ed States Unionidae. The Nautilus 10: 136–139. Lea, I. 1870. A synopsis of the family of Naı¨ades. Philadelphia, 4th edition, H. C. Lea, Philadelphia, 184 pp. APPENDIX 1 O’Brien, C. and J. Brim Box. 1999. Reproductive biology and SPECIMENS EXAMINED juvenile recruitment of the , Lampsilis subangulata (Bivalvia: Unionidae) in the Gulf Hamiota altilis Coastal Plain. American Midland Naturalist 142: 129–140. Parmalee, P. W. and A. E. Bogan. 1998. The Freshwater Mus- ALABAMA sels of Tennessee. The University of Tennessee Press, Knoxville, 328 pp. Alabama River: USNM 25948; Alabama River, near Clai- Rafinesque, C. S. 1819. Prodrome de 70 nouveaux generes borne: ANSP 56419 [Lectotype U. altilis Conrad, 1834], d’animaux de´couverts dans l’inte´rieur des Etats-Unis ANSP 56418; Big Swamp Creek, Macon Co.: USNM d’Ame´rique, Durant l’anne´e 1818. Journal de Physique, 361723; Jackson Shoals, Choccolocco Creek: ANSP de Chimie, d’Histoire Naturelle, et des Arts, Paris 88: 417–429. 103834, ANSP 103871; Beaver Creek: ANSP 103863; Rafinesque, C. S. 1820. Monographie des coquilles bivalves Coosa River: ANSP 41120, DMNH 130623; Coosa Riv- fluviatiles de la Rivie`re Ohio, contenant douze generes et er, Coosa Valley: ANSP 103771; Higgin’s Ferry, Coosa soixante-huit espe`ces. Annales ge´ne´rales des sciences Phy- River, Chilton Co.: USNM 218118; McGowen’s Ferry, siques, a Bruxelles 5: 287–322. near Wilsonville, Coosa River: USNM 521359; Weduska Reeve, L. 1865–1868. Monograph of the genus Unio. Con- Shoals, Coosa River: ANSP 48001, DMNH 075252, chologica Iconica. L. Reeve and Co., London. DMNH 150037, DMNH 150038, USNM 348970, Roe, K. J., P. D. Hartfield and C. Lydeard. 2001. Phylogeo- USNM 452169; Coosa River, [incomprehensible hand- graphic analysis of the threatened and endangered super- writing] Shoals, Shelby Co.: ANSP 341399; Shoal Creek, conglutinate-producing mussels of the genus Lampsilis Pine Glen Recreational Area, Cleburne Co.: MMNS (Bivalvia: Unionidae). Molecular Ecology 10: 2225–2234. 7743, MMNS 8084, MMNS 8085, UAUC 120, UAUC Simpson, C. T. 1892. Notes on the Unionidae of Florida and 121, UAUC 125; Yellowleaf Creek, Jumbo, Chilton Co.: the southeastern United States. Proceedings of the United States National Museum 15: 405–436, pls. 49–74. ANSP 89031; Cane Creek, West of CR 65, 2 mi. West Simpson, C. T. 1900a. Synopsis of the naiads or pearly fresh- CR 78 Jct., T15S, R11E, sec 3: UAUC 3292, MMNS water mussels. Proceedings of U.S. National Museum 22: 8081; Little Cane Creek, at CR 78, East of Edwardsville: ϳ 501–1044. UAUC 3293; Chewacla Creek at CR 22, 4 mi. East of Simpson, C. T. 1900b. New and unfigured Unionidae. Pro- Tuskeegee, Macon Co.: UAUC 246, UAUC 247, UAUC ceedings of the Academy of Natural Sciences of Philadel- 248; Uphappee Creek, 0.5 mi. upstream of Hwy. 29, phia 52: 74–86. Macon Co.: MMNS 8082; Cahaba River: USNM Swainson, W. 1840. A treatise on malacology or the natural 152026; Cahaba River: ANSP 126054; Little Cahaba classification of shells and shell-fish. London, 419 pp. River, 0.5 mi. below Cahaba Beach Rd. bridge, Jefferson Turgeon, D. D., J. F. Quinn, Jr., A. E. Bogan, E. V. Coan, F. Co.: UAUC 149; Coosa River, Weduska Shoals, Shelby G. Hochberg, W. G. Lyons, P. M. Mikkelsen, C. F. E. Co.: UF 3255; Coosa River at Fort William Shoals, Tal- Roper, G. Rosenberg, B. Roth, A. Scheltema, M. J. Swee- ladega Co.: UF 65420; Hurricane Creek, Cherokee Co.: ney, F. G. Thompson, M. Vecchione and J. D. Williams. UF 175098; Chewacla Creek, 8 mi. ESE of Tuskeegee 1988. Common and scientific names of aquatic inverte- brates from the United States and Canada: Mollusks, 2nd Co. Rd. 22, Macon Co.: UF 202249; Shoal Creek, St. edition. American Fisheries Society, Special Publication Clair Co.: UF 245989; Tuskeegee National Forest, Ma- 26, Bethesda, Maryland, 526 pp. con Co.: UF 266048; Cahaba River: UF 269576; Shoal U. S. Fish and Wildlife Service. 1994. Endangered and threat- Creek, St. Clair Co.: MMNS 8083. ened wildlife and plants, 50 CFR 17.11 and 17.12. Divi- sion of Endangered Species, U. S. Fish and Wildlife Ser- GEORGIA vice, Washington, D.C. U. S. Fish and Wildlife Service. 1998. Endangered and threat- Etowah River: USNM 84936 [Lectotype, U. doliaris ened wildlife and plants, Determination of endangered Lea, 1865]; Chattooga River: ANSP 89102; Chattooga K. J. Roe and P. D. Hartfield, 2005 Page 9

River, Chattooga Co.: USNM 59527; Chattanooga: UF 79076; Sipsey River at Sipsey Recreational Area, USNM 25711 [Holotype, U. gerhardtii, Lea, 1862]; Fish Bankhead N.F., Winston Co.: UF 79089; Borden’s Creek at Highway 278/ GA State Rt. 6, ϳ3.9 air mi. Creek, 1 mi., upstream of from Jct. Sipsey Fork, Sipsey West of Rockmart, Polk Co.: UAUC 538, UAUC 539; Wilderness, Bankhead National Forest, Winston Co.: Conasauga River at Tibbs Bridge Murray CR 109 (Whit- UF 79092, UF 79181; Sipsey Fork at N.F. Rd., 234, field CR100), Murray/Whitfield Co’s.: UAUC 515, Bankhead National Forest, Winston Co.: UF 79139; MMNS 8092; Conasauga River, Muskrat Midden, Tri- Limestone Creek, 6.3 mi WNW of Monroeville, Monroe togonia Shoals (CRM 46.70), Whitfield/Murray Co’s.: Co.: UF 197636; Blackwater Creek upstream from Har- UAUC 376; Conasauga River, south of state line, Murray ris bridge, Walker Co.: UF 266369; Sipsey River, 1.6 mi Co.: MMNS 8090; Conasauga River: DMNH 150124, N. of Pleasant Ridge, Greene Co.: UF 197671; North USNM 84937, USNM 348969; Etowah River: USNM River at Co. Hwy. 30, Fayette Co.: UF 197686; Sipsey 123202. River, 200 m. below Co. Hwy 23, Greene Co.: UF 197552; Sipsey River, 4–6 mi. below Co. Hwy. 2, Greene Co.: UF 197566; Sipsey River at CR 2, downstream of TENNESSEE boat ramp, Pickens Co.: UAUC 156; Tombigbee River: Conasauga River: DMNH 014683; Conasauga River, ANSP 126053; Elrod, Sipsey River, Tombigbee R.: Conasauga: ANSP 341305, ANSP 347949; Conasauga DMNH 146493; Lubbub Creek, at CR 24, 3.25 mi. River, Polk Co.: MMNS 8091. Northeast of Aliceville, Pickens Co.: UAUC 67; Tombig- bee River: USNM 159989; Lubbub Creek, 1.8 mi SSE Hamiota perovalis of Aliceville above Hwy 14: UF 197619, UF 197632; Sipsey River, 3.6 mi. W. of Jena downstream of CR 2, ALABAMA Greene Co.: UF 197697; Sipsey River near confluence with Carpenter’s Creek, Greene Co.: UF 197801; Sipsey Alabama River: ANSP 56416 [Lectotype U. perovalis River, 5.7 mi. NNE of Mantua, Greene Co.: UF 197857, Conrad, 1834], ANSP 333496; Alabama River, Clai- UF 197862; Sipsey River, near Elrod, Tuscaloosa Co.: borne: USNM 84938; Coosa River: ANSP 56415; Coosa UF 269559, UF 65301; Trussels Creek at CR 19 bridge, River, Gadsen: ANSP 126051; Coosa River, Talladega Greene Co.: MMNS 8087. Co.: ANSP 126048; Black Warrior River: ANSP 88483; Mulberry River [Fork, Black Warrior River]: ANSP MISSISSIPPI 88485; Rush Creek [Black Warrior River Dr.], FS Rt. 245, Winston Co.: MMNS 7745, MMNS 8088, UAUC Luxpalila Creek, near Columbus: USNM 84925 [Lec- 426; Flannigan Creek at FS Rd. 229, Lawrence Co.: totype, U. spillmanii Lea, 1861], USNM 123279; Colum- MMNS 7744, UAUC 423, UAUC 424, UAUC 425; bus, Lowndes Co.: UF 269560. Toadvine, Valley Creek, Black Warrior River Dr., Jeffer- son Co.: UF 65302, UF 65304; Forks of the Warrior Hamiota subangulata River, Walker Co.: UF 65305; North River, near Hagler’s Mill, Black Warrior Dr., Tuscaloosa Co.: UF 65306; Sip- ALABAMA sey Fork at N.F. 234, Bankhead National Forest, Win- ston Co.: UF 79069, UF 79072, UF 79082, UF 79085, Uchee Creek, Russell Co.: UAUC 116. UF 79136, UF 79137; Brushy Creek above Brushy Lake Recreational Area, Bankhead National Forest, Winston FLORIDA Co.: UF 79094; Capsey Creek, 50 mi. from Jct. with Brushy Creek, Bankhead N.F., Winston Co.: UF 79115; Chipola River, Look-Tremble Falls near Alpha, Calhoun Sipsey Fork at mouth of Hurricane Creek, Bankhead Co.: ANSP 175750; Chipola River, near Pole Bluff land- National Forest, Winston Co.: UAUC 95; Brown Creek, ing, Calhoun Co.: ANSP 175751; Chipola River: ANSP Bankhead National Forest, Winston Co.: UAUC 1774; 84324; Ochlockonee River: DMNH 150098; Ochlocko- Alabama: ANSP 126049; North River, near Samantha nee River, Leon Co.: USNM 149648 [Paratype, U. kirk- [Black Warrior River], Tuscaloosa Co.: UAUC 107; But- landianus S. H. Wright, 1897]; Ochlockonee River, Tal- tahatchee River, Hamilton: ANSP 100657, DMNH lahassee, Leon Co.: ANSP 156892, ANSP 341307; Och- 075231; Tuscaloosa Co. Alabama: DMNH 146496; Black lockonee River, 7 mi. west of Tallahassee: ANSP 157553; Warrior River, Jefferson Co.: UF 269609; Squaw Shoals, Ochlockonee River, 10 mi. west of Tallahassee, Leon Black Warrior River, Jefferson Co.: UF 65298, UF Co.: ANSP 159126; Ochlockonee River, 11 mi. north- 65299, UF 65300, UF 65303, UF 65307, UF 65429, UF west of Tallahassee: DMNH 119506; Ochlockonee Riv- 269518; Garden City, Mulberry Fork: UF 69207, UF er, 2 mi. west of Bloxham, Liberty Co.: ANSP 360553; 244558; Banks of Brushy Creek, at N.F. Rd. 255, Bank- Spring Creek, Marianna: ANSP 160210; State Rt. 167, head N.F, Winston Co.: UF 69269, UF 79177, UF 1 mi. north of Marianna, Chipola River, Jackson Co.: 79178, MMNS 7748, MMNS 8089; Sipsey Fork, 1 mile ANSP 349631; Spring Creek, Reynoldsville, Seminole N. of AL Hwy. 33 crossing, Winston Co.: UF 69279; Co.: UF 177; 1 mi. north Marianna, Chipola River: UF Brushy Creek at N.F. Rd., Bankhead N.F., Winston Co.: 390; 3.5 mi. east of Quincy, Little River: UF 415; ca. 2 Page 10 THE NAUTILUS, Vol. 119, No. 1 mi. east of Clarksville, Chipola River, Calhoun Co.: UF 74 bridge, Coweta/Fayette Co.: MMNS 8097; Ichaway- 418; Chipola River, 9.2 km ENE Kinard, 12.5 km NW nochaway Creek at GA Rt. 216, Baker Co.: MMNS Lewis, 16.4 km N. Ida, Calhoun Co.: MMNS 8099. 8098. Hamiota australis GEORGIA ALABAMA Chattahoochie River: ANSP 56477; Chattahoochie Riv- er, Columbus: USNM 85081 [Lectotype, U. subangula- Andrews fish trap, Pea River, Barbour Co.: UF 65309; tus Lea, 1840]; ANSP 126272; Cooleewahee Creek, 0.9 7 mi. east of Brundidge, Pea River, Pike Co.: UF mi. NE of Newton, Baker Co.: USNM 853746; Coolee- 123284; Bozemans landing, Conecuh River, near Cren- wahee Creek at GA Rt. 91, Baker Co.: MMNS 8095; shaw Co. line, Covington Co.: UF 65313; Lightwood Abram’s Inlet, Flint River: ANSP 190294; Mill Creek, Knot Creek, 1.6 mi. west of Opp, Covington Co.: Flint River, several mi. north of Albany: ANSP 267572; ANSP#; Little Patsaliga Creek: USNM 150473 [Holo- Kinchafoonee Creek at GA Rt. 41 crossing, just south of type, Lampsilis australis, Simpson, 1900]; West Fork Preston, Webster Co.: UAUC591; Kinchafoonee Creek Choctawhatchee River at Blue Spring State Park, Bar- at GA Rt. 49 bridge ϳ9 air mi. northeast of Dawson, bour Co.: UAUC 134, UAUC 511, UAUC 512, UAUC Terrell/Sumter Co’s.: UAUC 602, UAUC 603, UAUC 513, UAUC 514; Little Choctawhatchie Creek, near 604; Kinchafoonee Creek at GA Rt. 32, Lee Co., GA: Drian bridge, Houston Co.: UF 229532; Conecuh River MMNS 8096; Chickasawhatchee Creek at CR 130 on CR 28 ϳ1 mi. east of Goshen, Pike Co.: UAUC 510; bridge ϳ4 air mi. SW of Chickasawhatchee, Terrell Co.: Flat Creek at AL Hwy 153, near Flat Creek Church, UAUC 1753; Muckalee Creek at GA Rt. 195 bridge Geneva Co.: UAUC 547; Five Runs Creek, Conecuh N. ϳ3.5 air mi. Northeast of Leesburg, Lee Co.: UAUC F., Covington Co.: MMNS 8086. 312; Whitewater Creek on Morgan Mill Rd., Fayette Co.: UAUC 645; Ochlockonee River: DMNH 173390; FLORIDA Ochlockonee River, 7 mi. S. of Cairo: ANSP 194640, DMNH 048538, UF 412; Mimsville: ANSP 47892, Shoal River at Hwy 85, 1 mi. south of I-10 Jct. in Crest- DMNH 075151; Georgia: ANSP 126273; Calvary: ANSP view, Okaloosa Co.: UAUC 550, UAUC 551, UAUC 552, 47891; Ochlockonee River, Thomas/Grady Co’s. Georgia: UAUC 643, UAUC 644; Shoal Creek, ca. 1 mi. above MMNS 8101; Spring Creek at GA Rt. 84, Decatur Co.: U. S. Highway 90, Okaloosa Co.: UF 261852; Limestone MMNS 8094, MMNS 8100; Line Creek at GA Rt. 85/ Creek, Walton: MMNS 7746, MMNS 7747.